Revision of the Bark Beetle Genera Within the Former Cryphalini (curculionidae: Scolytinae)

Abstract Cryphalini Lindemann, 1877 are a speciose group of mostly miniscule beetles. The tribe Cryphalini is reviewed here which resulted in taxonomic and nomenclatural changes. This revision follows a recent molecular phylogenomic re-analysis focused on the tribe and related scolytine taxa. The analysis demonstrated that the tribe is polyphyletic, as found in other molecular phylogenies. To ensure monophyletic classification, we present a revision of the former tribe with two tribes resurrected, one described, and several genera transferred to other existing tribes. Additionally, extensive generic synonymy, and new combinations are presented. A key, photographs, and illustrations are provided to enable an accurate determination of genera. The revised Cryphalini contains only Cryphalus Erichson, 1836 (=Hypocryphalus Hopkins, 1915 syn. nov.; Margadillius Hopkins, 1915 syn. nov.). Coriacephilini Johnson trib. nov. contains only Coriacephilus Schedl, 1939. Ernoporini Nüsslin, 1911 stat. res. contains Eidophelus Eichhoff, 1876 (=Scolytogenes Eichhoff, 1878 syn. nov.; Ptilopodius Hopkins, 1915 syn. nov.; Ernoporicus Berger, 1917 syn. nov.; Cryphalogenes Wood, 1980 syn. nov.); ErnoporusThomson, 1859 (=Ernocladius Wood, 1980 syn. nov.; Allothenemus Bright and Torres, 2006 syn. nov.); Hemicryphalus Schedl, 1963; and Procryphalus Hopkins, 1915. Trypophloeini Nüsslin, 1911 stat. res. includes the genera Afrocosmoderes Johnson and Jordal gen. nov.; Atomothenemus Bright, 2019; Cosmoderes Eichhoff, 1878 (=Allernoporus Kurentsov, 1941 syn. nov.); Hypothenemus Westwood, 1834 (=Periocryphalus Wood, 1971 syn. nov.); Macrocryphalus Nobuchi, 1981 stat. res.; Microcosmoderes Johnson and Jordal gen. nov.; Microsomus Bright, 2019; Pygmaeoborus Bright, 2019; and Trypophloeus Fairmaire, 1864. Xyloterini LeConte, 1876 is maintained, containing Indocryphalus Eggers, 1939; Trypodendron Stephens, 1830 and Xyloterinus Swaine, 1918. Acorthylus Brèthes, 1922, Cryptocarenus Eggers, 1937, Neocryphus Nunberg, 1956, StegomerusWood, 1967, and Trypolepis Bright, 2019 are transferred to Corthylini LeConte, 1876. Stephanopodius Schedl, 1963 is transferred to Xyloctonini Eichhoff, 1878. As a consequence of generic synonymy, the following new or resurrected combinations are proposed: Cosmoderes euonymi (Kurentsov, 1941) comb. nov.; Cryphalus aciculatus (Schedl, 1939) comb. nov.; Cryphalus afiamalus (Schedl, 1951) comb. nov.; Cryphalus angustior Eggers, 1927 comb. res.; Cryphalus asper (Broun, 1881) comb. nov.; Cryphalus bakeri (Eggers, 1927) comb. nov.; Cryphalus basihirtus Beeson, 1929 comb. nov.; Cryphalus bidentatus (Browne, 1980) comb. nov.; Cryphalus brevior (Schedl, 1943) comb. nov.; Cryphalus carinatus (Browne, 1980) comb. nov.; Cryphalus confusus (Hopkins, 1915) comb. nov.; Cryphalus corpulentus (Schedl, 1942) comb. nov.; Cryphalus cylindripennis (Schedl, 1959) comb. nov.; Cryphalus cylindrus (Browne, 1950) comb. nov.; Cryphalus densepilosus (Schedl, 1942) comb. nov.; Cryphalus dilutus Eichhoff, 1878 comb. res.; Cryphalus discrepans (Schedl, 1965) comb. nov.; Cryphalus discretus Eichhoff, 1878 comb. res.; Cryphalus erythrinae (Hopkins, 1915) comb. nov.; Cryphalus fici (Browne, 1986) comb. nov.; Cryphalus glabratus (Schedl, 1959) comb. nov.; Cryphalus granulatus (Schedl, 1942) comb. nov.; Cryphalus imitans (Schedl, 1951) comb. nov.; Cryphalus interponens (Schedl, 1953) comb. nov.; Cryphalus kalambanganus (Schedl, 1943) comb. nov.; Cryphalus laevis (Browne, 1980) comb. nov.; Cryphalus laticollis (Browne, 1974) comb. nov.; Cryphalus longipennis (Browne, 1970) comb. nov.; Cryphalus longipilis (Browne, 1981) comb. nov.; Cryphalus magnus (Browne, 1984) comb. nov.; Cryphalus malayensis (Schedl, 1942) comb. nov.; Cryphalus mangiferae Stebbing, 1914 comb. res.; Cryphalus margadilaonis (Hopkins, 1915) comb. nov.; Cryphalus mindoroensis (Schedl, 1943) comb. nov.; Cryphalus minor (Schedl, 1943) comb. nov.; Cryphalus minutus (Hopkins, 1915) comb. nov.; Cryphalus mollis Schedl, 1955 comb. res.; Cryphalus moorei (Schedl, 1964) comb. nov.; Cryphalus nigrosetosus (Schedl, 1948) comb. nov.; Cryphalus nitidicollis (Schedl, 1975) comb. nov.; Cryphalus obscurus (Hopkins, 1915) comb. nov.; Cryphalus ovalicollis (Schedl, 1942) comb. nov.; Cryphalus papuanus (Schedl, 1973) comb. nov.; Cryphalus piliger (Schedl, 1975) comb. nov.; Cryphalus polynesiae (Schedl, 1979) comb. nov.; Cryphalus quadrituberculatus (Schedl, 1963) comb. nov.; Cryphalus reflexus (Browne, 1980) comb. nov.; Cryphalus robustus Eichhoff, 1872 comb. res.; Cryphalus rotundus (Hopkins, 1915) comb. nov.; Cryphalus sandakanensis Schedl, 1937 comb. res.; Cryphalus spathulatus (Schedl, 1938) comb. nov.; Cryphalus striatulus (Browne, 1978) comb. nov.; Cryphalus striatus (Hopkins, 1915) comb. nov.; Cryphalus sumatranus (Schedl, 1939) comb. nov.; Cryphalus triangularis (Schedl, 1975) comb. nov.; Cryphalus tutuilaensis (Schedl, 1951) comb. nov.; Eidophelus absonus (Schedl, 1975) comb. nov.; Eidophelus afer (Schedl, 1970) comb. nov.; Eidophelus africanus (Schedl, 1977) comb. nov.; Eidophelus aitutakii (Beaver and Maddison, 1990) comb. nov.; Eidophelus alniphagus (Nobuchi, 1975) comb. nov.; Eidophelus alternans (Schedl, 1975) comb. nov.; Eidophelus amanicus (Eggers, 1919) comb. nov.; Eidophelus ankius (Schedl, 1979) comb. nov.; Eidophelus apicalis (Schedl, 1971) comb. nov.; Eidophelus approximatus (Schedl, 1975) comb. nov.; Eidophelus aspericollis (Eichhoff, 1878) comb. nov.; Eidophelus ater (Eggers, 1923) comb. nov.; Eidophelus australis (Schedl, 1942) comb. nov.; Eidophelus badius (Nobuchi, 1975) comb. nov.; Eidophelus bambusae (Browne, 1983) comb. nov.; Eidophelus bangensis (Eggers, 1927) comb. nov.; Eidophelus basilaris (Wood, 1960) comb. nov.; Eidophelus birosimensis (Murayama, 1958) comb. nov.; Eidophelus braderi (Browne, 1965) comb. nov.; Eidophelus brimblecombei (Schedl, 1972) comb. nov.; Eidophelus buruensis (Eggers, 1926) comb. nov.; Eidophelus camelliae (Nobuchi, 1975) comb. nov.; Eidophelus candidus (Nobuchi, 1975) comb. nov.; Eidophelus capucinus (Schedl, 1971) comb. nov.; Eidophelus caucasicus (Lindemann, 1877) comb. nov.; Eidophelus ceylonicus (Schedl, 1959) comb. nov.; Eidophelus cicatricosus (Schedl, 1942) comb. nov.; Eidophelus coccotrypanoides (Schedl, 1939) comb. nov.; Eidophelus communis (Schaufuss, 1891) comb. nov.; Eidophelus confragosus (Sampson, 1914) comb. nov.; Eidophelus corni (Kurentsov, 1941) comb. nov.; Eidophelus corpulentus (Schedl, 1965) comb. nov.; Eidophelus corrugatus (Schedl, 1950) comb. nov.; Eidophelus creber (Schedl, 1975) comb. nov.; Eidophelus crenatus (Sampson, 1914) comb. nov.; Eidophelus cylindricus (Schedl, 1959) comb. nov.; Eidophelus darwini (Eichhoff, 1878) comb. nov.; Eidophelus devius (Schedl, 1975) comb. nov.; Eidophelus dubiosus (Wood, 1960) comb. nov.; Eidophelus eggersi (Schedl, 1962) comb. nov.; Eidophelus euphorbiae (Wood, 1980) comb. nov.; Eidophelus excellens (Schedl, 1979) comb. nov.; Eidophelus exiguus (Wood, 1980) comb. nov.; Eidophelus exilis (Yin, 2001) comb. nov.; Eidophelus eximius (Schedl, 1942) comb. nov.; Eidophelus expers (Blandford, 1894) comb. nov.; Eidophelus fagi (Fabricius, 1798) comb. nov.; Eidophelus fijianus (Schedl, 1950) comb. nov.; Eidophelus formosanus (Browne, 1981) comb. nov.; Eidophelus fugax (Schedl, 1975) comb. nov.; Eidophelus fujisanus (Nobuchi, 1975) comb. nov.; Eidophelus fulgens (Schedl, 1975) comb. nov.; Eidophelus fulgidus (Schedl, 1975) comb. nov.; Eidophelus fulvipennis (Nobuchi, 1975) comb. nov.; Eidophelus ghanaensis (Schedl, 1977) comb. nov.; Eidophelus glabratus (Yin, 2001) comb. nov.; Eidophelus gracilis (Schedl, 1950) comb. nov.; Eidophelus granulatus (Wood, 1960) comb. nov.; Eidophelus grobleri (Schedl, 1962) comb. nov.; Eidophelus hirtus (Wood, 1974) comb. nov.; Eidophelus hobohmi (Schedl, 1955) comb. nov.; Eidophelus hylesinopsis (Schedl, 1975) comb. nov.; Eidophelus incultus (Yin, 2001) comb. nov.; Eidophelus indicus ((Wood, 1989)) comb. nov.; Eidophelus insularis (Nobuchi, 1975) comb. nov.; Eidophelus insularum (Krivolutskaya, 1968) comb. nov.; Eidophelus jalappae (Letzner, 1849) comb. nov.; Eidophelus javanus (Schedl, 1942) comb. nov.; Eidophelus kanawhae (Hopkins, 1915) comb. nov.; Eidophelus landolphiae (Schedl, 1961) comb. nov.; Eidophelus leprosulus (Browne, 1974) comb. nov.; Eidophelus longipennis (Eggers, 1936) comb. nov.; Eidophelus magnocularis (Yin, 2001) comb. nov.; Eidophelus marquesanus (Beeson, 1935) comb. nov.; Eidophelus mauritianus (Schedl, 1965) comb. nov.; Eidophelus micans (Eggers, 1927) comb. nov.; Eidophelus minor (Eggers, 1927) comb. nov.; Eidophelus minutissimus (Schedl, 1943) comb. nov.; Eidophelus mus (Schedl, 1975) comb. nov.; Eidophelus nanulus (Wood, 1960) comb. nov.; Eidophelus nigellatus (Schedl, 1950) comb. nov.; Eidophelus nubilus (Wood, 1960) comb. nov.; Eidophelus ocularis (Schedl, 1965) comb. nov.; Eidophelus onyanganus (Schedl, 1941) comb. nov.; Eidophelus opacus (Schedl, 1959) comb. nov.; Eidophelus pacificus (Schedl, 1941) comb. nov.; Eidophelus papuanus (Schedl, 1974) comb. nov.; Eidophelus papuensis ((Wood, 1989)) comb. nov.; Eidophelus paradoxus (Wood, 1992) comb. nov.; Eidophelus parvus (Hopkins, 1915) comb. nov.; Eidophelus pityophthorinus (Schedl, 1943) comb. nov.; Eidophelus pleiocarpae (Schedl, 1957) comb. nov.; Eidophelus polisquamosus (Yin, 2001) comb. nov.; Eidophelus praeda (Browne, 1978) comb. nov.; Eidophelus puerarae (Choo and Woo, 1989) comb. nov.; Eidophelus pumilionides (Schedl, 1977) comb. nov.; Eidophelus pumilus (Wood, 1960) comb. nov.; Eidophelus punctatulus (Nobuchi, 1976) comb. nov.; Eidophelus punctatus (Schedl, 1951) comb. nov.; Eidophelus puncticollis (Schedl, 1950) comb. nov.; Eidophelus pygmaeolus (Schedl, 1971) comb. nov.; Eidophelus quadridens (Browne, 1983) comb. nov.; Eidophelus ramosus (Beeson, 1935) comb. nov.; Eidophelus robustus (Schedl, 1955) comb. nov.; Eidophelus rugosus (Schedl, 1943) comb. nov.; Eidophelus rusticus (Wood, 19